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1.  Phenols in Leaves and Bark of Fagus sylvatica as Determinants of Insect Occurrences 
Beech forests play an important role in temperate and north Mediterranean ecosystems in Greece since they occupy infertile montane soils. In the last glacial maximum, Fagus sylvatica (beech) was confined to Southern Europe where it was dominant and in the last thousand years has expanded its range to dominate central Europe. We sampled four different beech forest types. We found 298 insect species associated with beech trees and dead beech wood. While F. sylvatica and Quercus (oak) are confamilial, there are great differences in richness of the associated entomofauna. Insect species that inhabit beech forests are less than one fifth of those species living in oak dominated forests despite the fact that beech is the most abundant central and north European tree. There is a distinct paucity of monophagous species on beech trees and most insect species are shared between co-occurring deciduous tree species and beech. This lack of species is attributed to the vegetation history and secondary plant chemistry. Bark and leaf biophenols from beech indicate that differences in plant secondary metabolites may be responsible for the differences in the richness of entomofauna in communities dominated by beech and other deciduous trees.
PMCID: PMC3116155  PMID: 21686149
Fagus sylvatica; entomofauna; phenolics; secondary plant chemistry; insect traps; clustering; discriminant analysis
2.  Does mixing of beech (Fagus sylvatica) and spruce (Picea abies) litter hasten decomposition? 
Plant and soil  2014;377(1-2):217-234.
Background and aims
It is of practical relevance to know how much beech must be admixed to pure spruce stands in order to increase litter decomposition and associated nutrient cycling, since the formation of thick organic layers is commonly ascribed to the recalcitrance of spruce needles. We addressed the impact of tree species mixture within forest stands and within litter on mass loss and nutritional release from litter.
Litter decomposition was measured in three adjacent stands of pure spruce (Picea abies), mixed beech-spruce and pure beech (Fagus sylvatica) on a nutrient-rich site and a nutrient-poor site over a 2-year period using litterbags which were filled with five different mixtures of beech and spruce litter.
Mass loss of beech litter was not higher than mass loss of spruce litter. Decay was primarily affected by tree species composition of the incubation stand and was faster in (mixed) beech forests stands than in spruce forests, while the influence of litter species and their mixtures on decay rates was small. Net transfers of nutrients between the two litter species (direct effects) in the mixed bags were minimal, since initial beech and spruce litter did not have different litter quality. However, in a few cases indirect effects (e.g., changing decomposer abundance and activity) caused non-additive patterns for the totals within the mixed bags, hastening decomposition within the first year.
Greater accumulation of litter in spruce compared to beech stands is not a consequence of the inherent recalcitrance of needles. Adverse environmental conditions in spruce stands retard decomposition. Indirect effects on decomposition caused by stand mixture are not mimicked by litter mixtures within mesh bags.
PMCID: PMC3987168  PMID: 24744450
Decomposition; Fagus sylvatica; Litter quality; Litterbag; Mixing effects; Picea abies
3.  Impact of elevated atmospheric O3 on the actinobacterial community structure and function in the rhizosphere of European beech (Fagus sylvatica L.) 
Many bacteria belonging to the phylum of Actinobacteria are known as antagonists against phytpathogenic microbes. This study aimed to analyze the effect of ozone on the actinobacterial community of the rhizosphere of four years old European beech (Fagus sylvatica L.) trees during different time points of the vegetation period. Effects of ozone on the total community structure of Actinobacteria were studied based on the analysis of 16S rRNA gene amplicons. In addition effects of the ozone treatment on the diversity of potential biocontrol active Actionobacteria being able to produce antibiotics were characterized by using the type II polyketide synthases (PKS) genes as marker. Season as well as ozone treatments had a significant effect on parts of the actinobacterial rhizosphere community of European beech. However on the basis of the performed analysis, the diversity of Actinobacteria possessing type II PKS genes is neither affected by seasonal changes nor by the ozone treatments, indicating no influence of the investigated treatments on the biocontrol active part of the actinobacterial community.
PMCID: PMC3920289  PMID: 24575080
European beech; Actinobacteria; biocontrol; polyketide synthase gene; ozone
4.  Drought-Adaptation Potential in Fagus sylvatica: Linking Moisture Availability with Genetic Diversity and Dendrochronology 
PLoS ONE  2012;7(3):e33636.
Microevolution is essential for species persistence especially under anticipated climate change scenarios. Species distribution projection models suggested that the dominant tree species of lowland forests in Switzerland, European beech (Fagus sylvatica L.), might disappear from most areas due to expected longer dry periods. However, if genotypes at the moisture boundary of the species climatic envelope are adapted to lower moisture availability, they can serve as seed source for the continuation of beech forests under changing climates.
Methodology/Principal Findings
With an AFLP genome scan approach, we studied neutral and potentially adaptive genetic variation in Fagus sylvatica in three regions containing a dry and a mesic site each (nind. = 241, nmarkers = 517). We linked this dataset with dendrochronological growth measures and local moisture availabilities based on precipitation and soil characteristics. Genetic diversity decreased slightly at dry sites. Overall genetic differentiation was low (Fst = 0.028) and Bayesian cluster analysis grouped all populations together suggesting high (historical) gene flow. The Bayesian outlier analyses indicated 13 markers with three markers differing between all dry and mesic sites and the others between the contrasting sites within individual regions. A total of 41 markers, including seven outlier loci, changed their frequency with local moisture availability. Tree height and median basal growth increments were reduced at dry sites, but marker presence/absence was not related to dendrochronological characteristics.
Conclusion and Their Significance
The outlier alleles and the makers with changing frequencies in relation to moisture availability indicate microevolutionary processes occurring within short geographic distances. The general genetic similarity among sites suggests that ‘preadaptive’ genes can easily spread across the landscape. Yet, due to the long live span of trees, fostering saplings originating from dry sites and grown within mesic sites might increase resistance of beech forests during the anticipated longer dry periods.
PMCID: PMC3308988  PMID: 22448260
5.  Leaf litter decomposition in temperate deciduous forest stands with a decreasing fraction of beech (Fagus sylvatica) 
Oecologia  2010;164(4):1083-1094.
We hypothesised that the decomposition rates of leaf litter will increase along a gradient of decreasing fraction of the European beech (Fagus sylvatica) and increasing tree species diversity in the generally beech-dominated Central European temperate deciduous forests due to an increase in litter quality. We studied the decomposition of leaf litter including its lignin fraction in monospecific (pure beech) stands and in stands with up to five tree genera (Acer spp., Carpinus betulus, Fagus sylvatica, Fraxinus excelsior, Tilia spp.) using a litterbag approach. Litter and lignin decomposition was more rapid in stand-representative litter from multispecific stands than in litter from pure beech stands. Except for beech litter, the decomposition rates of species-specific tree litter did not differ significantly among the stand types, but were most rapid in Fraxinus excelsior and slowest in beech in an interspecific comparison. Pairwise comparisons of the decomposition of beech litter with litter of the other tree species (except for Acerplatanoides) revealed a “home field advantage” of up to 20% (more rapid litter decomposition in stands with a high fraction of its own species than in stands with a different tree species composition). Decomposition of stand-representative litter mixtures displayed additive characteristics, not significantly more rapid than predicted by the decomposition of litter from the individual tree species. Leaf litter decomposition rates were positively correlated with the initial N and Ca concentrations of the litter, and negatively with the initial C:N, C:P and lignin:N ratios. The results support our hypothesis that the overall decomposition rates are mainly influenced by the chemical composition of the individual litter species. Thus, the fraction of individual tree species in the species composition seems to be more important for the litter decomposition rates than tree species diversity itself.
PMCID: PMC2981742  PMID: 20596729
Biodiversity; Central Europe; Element ratio; Litter degradation; Litterbag
6.  Sapling herbivory, invertebrate herbivores and predators across a natural tree diversity gradient in Germany’s largest connected deciduous forest 
Oecologia  2009;160(2):279-288.
Tree species-rich forests are hypothesised to be less susceptible to insect herbivores, but so far herbivory–diversity relationships have rarely been tested for tree saplings, and no such study has been published for deciduous forests in Central Europe. We expected that diverse tree communities reduce the probability of detection of host plants and increase abundance of predators, thereby reducing herbivory. We examined levels of herbivory suffered by beech (Fagus sylvatica L.) and maple saplings (Acer pseudoplatanus L. and Acer platanoides L.) across a tree species diversity gradient within Germany’s largest remaining deciduous forest area, and investigated whether simple beech or mixed stands were less prone to damage caused by herbivorous insects. Leaf area loss and the frequency of galls and mines were recorded for 1,040 saplings (>13,000 leaves) in June and August 2006. In addition, relative abundance of predators was assessed to test for potential top-down control. Leaf area loss was generally higher in the two species of maple compared to beech saplings, while only beech showed a decline in damage caused by leaf-chewing herbivores across the tree diversity gradient. No significant patterns were found for galls and mines. Relative abundance of predators on beech showed a seasonal response and increased on species-rich plots in June, suggesting higher biological control. We conclude that, in temperate deciduous forests, herbivory–tree diversity relationships are significant, but are tree species-dependent with bottom-up and top-down control as possible mechanisms. In contrast to maple, beech profits from growing in a neighbourhood of higher tree richness, which implies that species identity effects may be of greater importance than tree diversity effects per se. Hence, herbivory on beech appeared to be mediated bottom-up by resource concentration in the sampled forest stands, as well as regulated top-down through biocontrol by natural enemies.
Electronic supplementary material
The online version of this article (doi:10.1007/s00442-009-1304-2) contains supplementary material, which is available to authorised users.
PMCID: PMC3085765  PMID: 19238448
Diversity-functioning relationships; Leaf damage; Mines; Multitrophic interactions; Plant–animal interactions
7.  Establishment success of sooty beech scale insects, Ultracoelostoma sp., on different host tree species in New Zealand 
The sooty beech scale insect (Ultracoelostoma sp.) (Hemiptera: Margarodidae) exhibits a highly patchy distribution at local and regional scales. A major factor driving this common distributional phenomenon in other phloem-feeding insects is aggregation and local adaptation. The aim of this study was to determine if Ultracoelostoma was locally adapted to its natal host trees, by contrasting the establishment rates of first instar “crawlers” in reciprocal transfers to natal versus novel hosts. Although there are two closely-related species of sooty beech scale insect, the morphological characters of crawlers in this study were intermediate between those of U. assimile and U. brittini. However, all of the voucher specimens examined had consistent morphology, indicating that they belong to one species which we refer to as Ultracoelostoma sp. Reciprocal transfers of crawlers were carried out between individual red beech (Nothofagus fusca), as well as between mountain beech (N. solandri) and red beech trees, to ascertain if insects had become locally adapted to their individual host tree or to host species. In total, 480 crawlers were placed in enclosures on their natal and novel host trees, of which only 32 (6.7 %) became established. No evidence for local adaptation, either to individual host trees or to host tree species, was found. There was also no difference in crawler establishment between natal and novel hosts. However, crawlers originating from mountain beech trees had significantly higher establishment rates on both natal mountain beech and novel red beech hosts, than did crawlers originating from red beech trees. The superior ability of mountain beech crawlers to become established, even on novel red beech trees, suggests that scale insects on mountain beech trees have higher individual fitness (possibly due to maternal effects mediated by differences in host nutritional quality, defensive compounds or growth rate). This increased fitness may result in crawlers being better provisioned to search for appropriate establishment sites. The results of this study indicate that beech scale insects perform better on mountain beech at this site, although crawlers did not preferentially establish on mountain beech.
PMCID: PMC2990322  PMID: 19537979
honeydew; host preference; local adaptation; Ultracoelostoma assimile; Ultracoelostoma brittini; Nothofagus fusca; Nothofagus solandri
8.  Model-based analysis of avoidance of ozone stress by stomatal closure in Siebold's beech (Fagus crenata) 
Annals of Botany  2013;112(6):1149-1158.
Background and Aims
Resistance of plants to ozone stress can be classified as either avoidance or tolerance. Avoidance of ozone stress may be explained by decreased stomatal conductance during ozone exposure because stomata are the principal interface for entry of ozone into plants. In this study, a coupled photosynthesis–stomatal model was modified to test whether the presence of ozone can induce avoidance of ozone stress by stomatal closure.
The response of Siebold's beech (Fagus crenata), a representative deciduous tree species, to ozone was studied in a free-air ozone exposure experiment in Japan. Photosynthesis and stomatal conductance were measured under ambient and elevated ozone. An optimization model of stomata involving water, CO2 and ozone flux was tested using the leaf gas exchange data.
Key Results
The data suggest that there are two phases in the avoidance of ozone stress via stomatal closure for Siebold's beech: (1) in early summer ozone influx is efficiently limited by a reduction in stomatal conductance, without any clear effect on photosynthetic capacity; and (2) in late summer and autumn the efficiency of ozone stress avoidance was decreased because the decrease in stomatal conductance was small and accompanied by an ozone-induced decline of photosynthetic capacity.
Ozone-induced stomatal closure in Siebold's beech during early summer reduces ozone influx and allows the maximum photosynthetic capacity to be reached, but is not sufficient in older leaves to protect the photosynthetic system.
PMCID: PMC3783231  PMID: 23904447
Tropospheric ozone; stomatal conductance; stomatal closure; stress avoidance; photosynthesis–stomatal model; Siebold's beech; Fagus crenata
9.  Girdling Affects Ectomycorrhizal Fungal (EMF) Diversity and Reveals Functional Differences in EMF Community Composition in a Beech Forest▿ † 
The relationships between plant carbon resources, soil carbon and nitrogen content, and ectomycorrhizal fungal (EMF) diversity in a monospecific, old-growth beech (Fagus sylvatica) forest were investigated by manipulating carbon flux by girdling. We hypothesized that disruption of the carbon supply would not affect diversity and EMF species numbers if EM fungi can be supplied by plant internal carbohydrate resources or would result in selective disappearance of EMF taxa because of differences in carbon demand of different fungi. Tree carbohydrate status, root demography, EMF colonization, and EMF taxon abundance were measured repeatedly during 1 year after girdling. Girdling did not affect root colonization but decreased EMF species richness of an estimated 79 to 90 taxa to about 40 taxa. Cenococcum geophilum, Lactarius blennius, and Tomentella lapida were dominant, colonizing about 70% of the root tips, and remained unaffected by girdling. Mainly cryptic EMF species disappeared. Therefore, the Shannon-Wiener index (H′) decreased but evenness was unaffected. H′ was positively correlated with glucose, fructose, and starch concentrations of fine roots and also with the ratio of dissolved organic carbon to dissolved organic nitrogen (DOC/DON), suggesting that both H′ and DOC/DON were governed by changes in belowground carbon allocation. Our results suggest that beech maintains numerous rare EMF species by recent photosynthate. These EM fungi may constitute biological insurance for adaptation to changing environmental conditions. The preservation of taxa previously not known to colonize beech may, thus, form an important reservoir for future forest development.
PMCID: PMC2837996  PMID: 20097809
10.  Physiological Responses of Beech and Sessile Oak in a Natural Mixed Stand During a Dry Summer 
Annals of Botany  2002;89(6):723-730.
Responses of CO2 assimilation and stomatal conductance to decreasing leaf water potential, and to environmental factors, were analysed in a mixed natural stand of sessile oak (Quercus petraea ssp. medwediewii) and beech (Fagus sylvatica L.) in Greece during the exceptionally dry summer of 1998. Seasonal courses of leaf water potential were similar for both species, whereas mean net photosynthesis and stomatal conductance were always higher in sessile oak than in beech. The relationship between net photosynthesis and stomatal conductance was strong for both species. Sessile oak had high rates of photosynthesis even under very low leaf water potentials and high air temperatures, whereas the photosynthetic rate of beech decreased at low water potentials. Diurnal patterns were similar in both species but sessile oak had higher rates of CO2 assimilation than beech. Our results indicate that sessile oak is more tolerant of drought than beech, due, in part, to its maintenance of photosynthesis at low water potential.
PMCID: PMC4233840  PMID: 12102528
Sessile oak; Quercus petraea ssp.; medwediewii; beech; Fagus sylvatica L.; CO2 assimilation; stomatal conductance; leaf water potential; drought
11.  Gap Dynamics and Structure of Two Old-Growth Beech Forest Remnants in Slovenia 
PLoS ONE  2013;8(1):e52641.
Due to a long history of intensive forest exploitation, few European beech (Fagus sylvatica L.) old-growth forests have been preserved in Europe.
Material and Methods
We studied two beech forest reserves in southern Slovenia. We examined the structural characteristics of the two forest reserves based on data from sample plots and complete inventory obtained from four previous forest management plans. To gain a better understanding of disturbance dynamics, we used aerial imagery to study the characteristics of canopy gaps over an 11-year period in the Kopa forest reserve and a 20-year period in the Gorjanci forest reserve.
The results suggest that these forests are structurally heterogeneous over small spatial scales. Gap size analysis showed that gaps smaller than 500 m2 are the dominant driving force of stand development. The percentage of forest area in canopy gaps ranged from 3.2 to 4.5% in the Kopa forest reserve and from 9.1 to 10.6% in the Gorjanci forest reserve. These forests exhibit relatively high annual rates of coverage by newly established (0.15 and 0.25%) and closed (0.08 and 0.16%) canopy gaps. New gap formation is dependant on senescent trees located throughout the reserve.
We conclude that these stands are not even-sized, but rather unevenly structured. This is due to the fact that the disturbance regime is characterized by low intensity, small-scale disturbances.
PMCID: PMC3538742  PMID: 23308115
12.  A Dynamical Model of Environmental Effects on Allocation to Carbon-based Secondary Compounds in Juvenile Trees 
Annals of Botany  2007;101(8):1089-1098.
Background and Aims
Patterns and variations in concentration of carbon-based secondary compounds in plant tissues have been explained by means of different complementary and, in some cases, contradictory plant defence hypotheses for more than 20 years. These hypotheses are conceptual models which consider environmental impacts on plant internal demands. In the present study, a mathematical model is presented, which converts and integrates the concepts of the ‘Growth–Differentiation Balance’ hypothesis and the ‘Protein Competition’ model into a dynamic plant growth model, that was tested with concentration data of polyphenols in leaves of juvenile apple, beech and spruce trees. The modelling approach is part of the plant growth model PLATHO that considers simultaneously different environmental impacts on the most important physiological processes of plants.
The modelling approach for plant internal resource allocation is based on a priority scheme assuming that growth processes have priority over allocation to secondary compounds and that growth-related metabolism is more strongly affected by nitrogen deficiency than defence-related secondary metabolism.
Key Results
It is shown that the model can reproduce the effect of nitrogen fertilization on allocation patterns in apple trees and the effects of elevated CO2 and competition in juvenile beech and spruce trees. The analysis of model behaviour reveals that large fluctuations in plant internal availability of carbon and nitrogen are possible within a single vegetation period. Furthermore, the model displays a non-linear allocation behaviour to carbon-based secondary compounds.
The simulation results corroborate the underlying assumptions of the presented modelling approach for resource partitioning between growth-related primary metabolism and defence-related secondary metabolism. Thus, the dynamical modelling approach, which considers variable source and sink strengths of plant internal resources within different phenological growth stages, presents a successful translation of existing concepts into a dynamic mathematical model.
PMCID: PMC2710266  PMID: 17693454
Plant growth; carbon-based secondary compounds; plant defence hypotheses; simulation model; phenolic allocation; nitrogen; carbon dioxide; Malus domestica; Fagus sylvatica; Picea abies
13.  Space sequestration below ground in old-growth spruce-beech forests—signs for facilitation? 
Scientists are currently debating the effects of mixing tree species for the complementary resource acquisition in forest ecosystems. In four unmanaged old-growth spruce-beech forests in strict nature reserves in southern Sweden and northern Germany we assessed forest structure and fine rooting profiles and traits (≤2 mm) by fine root sampling and the analysis of fine root morphology and biomass. These studies were conducted in selected tree groups with four different interspecific competition perspectives: (1) spruce as a central tree, (2) spruce as competitor, (3) beech as a central tree, and (4) beech as competitor. Mean values of life fine root attributes like biomass (FRB), length (FRL), and root area index (RAI) were significantly lower for spruce than for beech in mixed stands. Vertical profiles of fine root attributes adjusted to one unit of basal area (BA) exhibited partial root system stratification when central beech is growing with spruce competitors. In this constellation, beech was able to raise its specific root length (SRL) and therefore soil exploration efficiency in the subsoil, while increasing root biomass partitioning into deeper soil layers. According to relative values of fine root attributes (rFRA), asymmetric below-ground competition was observed favoring beech over spruce, in particular when central beech trees are admixed with spruce competitors. We conclude that beech fine rooting is facilitated in the presence of spruce by lowering competitive pressure compared to intraspecific competition whereas the competitive pressure for spruce is increased by beech admixture. Our findings underline the need of spatially differentiated approaches to assess interspecific competition below ground. Single-tree approaches and simulations of below-ground competition are required to focus rather on microsites populated by tree specimens as the basic spatial study area.
PMCID: PMC3747362  PMID: 24009616
Fagus sylvatica; Picea abies; root system stratification; fine root biomass (FRB); fine root length (FRL); fine root surface area index (RAI); specific root length (SRL); specific root surface area (SRA)
14.  Different Atmospheric Methane-Oxidizing Communities in European Beech and Norway Spruce Soils▿ †  
Applied and Environmental Microbiology  2010;76(10):3228-3235.
Norway spruce (Picea abies) forests exhibit lower annual atmospheric methane consumption rates than do European beech (Fagus sylvatica) forests. In the current study, pmoA (encoding a subunit of membrane-bound CH4 monooxygenase) genes from three temperate forest ecosystems with both beech and spruce stands were analyzed to assess the potential effect of tree species on methanotrophic communities. A pmoA sequence difference of 7% at the derived protein level correlated with the species-level distance cutoff value of 3% based on the 16S rRNA gene. Applying this distance cutoff, higher numbers of species-level pmoA genotypes were detected in beech than in spruce soil samples, all affiliating with upland soil cluster α (USCα). Additionally, two deep-branching genotypes (named 6 and 7) were present in various soil samples not affiliating with pmoA or amoA. Abundance of USCα pmoA genes was higher in beech soils and reached up to (1.2 ± 0.2) × 108 pmoA genes per g of dry weight. Calculated atmospheric methane oxidation rates per cell yielded the same trend. However, these values were below the theoretical threshold necessary for facilitating cell maintenance, suggesting that USCα species might require alternative carbon or energy sources to thrive in forest soils. These collective results indicate that the methanotrophic diversity and abundance in spruce soils are lower than those of beech soils, suggesting that tree species-related factors might influence the in situ activity of methanotrophs.
PMCID: PMC2869149  PMID: 20348309
15.  Spatial Patterns of Ectomycorrhizal Assemblages in a Monospecific Forest in Relation to Host Tree Genotype 
Ectomycorrhizas (EcM) are important for soil exploration and thereby may shape belowground interactions of roots. We investigated the composition and spatial structures of EcM assemblages in relation to host genotype in an old-growth, monospecific beech (Fagus sylvatica) forest. We hypothesized that neighboring roots of different beech individuals are colonized by similar EcM assemblages if host genotype had no influence on the fungal colonization and that the similarity would decrease with increasing distance of the sampling points. The alternative was that the EcM species showed preferences for distinct beech genotypes resulting in intraspecific variation of EcM-host assemblages. EcM species identities, abundance and exploration type as well as the genotypes of the colonized roots were determined in each sampling unit of a 1 L soil core (r = 0.04 m, depth 0.2 m). The Morisita-Horn similarity indices (MHSI) based on EcM species abundance and multiple community comparisons were calculated. No pronounced variation of MHSI with increasing distances of the sampling points within a plot was found, but variations between plots. Very high similarities and no between plot variation were found for MHSI based on EcM exploration types suggesting homogenous soil foraging in this ecosystem. The EcM community on different root genotypes in the same soil core exhibited high similarity, whereas the EcM communities on the root of the same tree genotype in different soil cores were significantly dissimilar. This finding suggests that spatial structuring of EcM assemblages occurs within the root system of an individual. This may constitute a novel, yet unknown mechanism ensuring colonization by a diverse EcM community of the roots of a given host individual.
PMCID: PMC3633777  PMID: 23630537
belowground interactions; community ecology; ectomycorrhiza; deciduous forest; intraspecific variation; interspecific variation
16.  Leaf traits, shoot growth and seed production in mature Fagus sylvatica trees after 8 years of CO2 enrichment 
Annals of Botany  2011;107(8):1405-1411.
Background and Aims
Masting, i.e. synchronous but highly variable interannual seed production, is a strong sink for carbon and nutrients. It may, therefore, compete with vegetative growth. It is currently unknown whether increased atmospheric CO2 concentrations will affect the carbon balance (or that of other nutrients) between reproduction and vegetative growth of forest species. In this study, reproduction and vegetative growth of shoots of mature beech (Fagus sylvatica) trees grown at ambient and elevated atmospheric CO2 concentrations were quantified. It was hypothesized that within a shoot, fruiting has a negative effect on vegetative growth, and that this effect is ameliorated at increased CO2 concentrations.
Reproduction and its competition with leaf and shoot production were examined during two masting events (in 2007 and 2009) in F. sylvatica trees that had been exposed to either ambient or elevated CO2 concentrations (530 µmol mol−1) for eight consecutive years, between 2000 and 2008.
Key Results
The number of leaves per shoot and the length of terminal shoots was smaller or shorter in the two masting years compared with the one non-masting year (2008) investigated, but they were unaffected by elevated CO2 concentrations. The dry mass of terminal shoots was approx. 2-fold lower in the masting year (2007) than in the non-masting year in trees growing at ambient CO2 concentrations, but this decline was not observed in trees exposed to elevated CO2 concentrations. In both the CO2 treatments, fruiting significantly decreased nitrogen concentration by 25 % in leaves and xylem tissue of 1- to 3-year-old branches in 2009.
Our findings indicate that there is competition for resources between reproduction and shoot growth. Elevated CO2 concentrations reduced this competition, indicating effects on the balance of resource allocation between reproduction and vegetative growth in shoots with rising atmospheric CO2 concentrations.
PMCID: PMC3101148  PMID: 21493641
Beech; carbon autonomy; CO2 enrichment; Fagus sylvatica; mast seeding; nitrogen; resource allocation; trade-off; vegetative growth
17.  Introduction to Distribution and Ecology of Sterile Conks of Inonotus obliquus 
Mycobiology  2008;36(4):199-202.
Inonotus obliquus is a fungus that causes white heart rot on several broad-leaved species. This fungus forms typical charcoal-black, sterile conks (chaga) or cinder conks on infected stems of the birche (Betula spp). The dark brown pulp of the sterile conk is formed by a pure mycelial mass of fungus. Chaga are a folk remedy in Russia, reflecting the circumboreal distribution of I. obliquus in boreal forest ecosystems on Betula spp. and in meridional mountain forests on beech (Fagus spp.) in Russia, Scandinavia, Central Europe, and Eastern Europe. Distribution at lower latitudes in Western and Southern Europe, Northern America, Asia, Japan, and Korea is rare. Infected trees grow for many years without several symptoms of decline. The infection can penetrate through stem injuries with exterior sterile conks developing later. In the Czech Republic, cinder conk is found on birches inhabiting peat bogs and in mountain areas with a colder and more humid climate, although it is widespread in other broad leaved species over the Czech Republic. The most common hosts are B. pendula, B. pubescens, B. carpatica, and F. sylvatica. Less frequent hosts include Acer campestre, Acer pseudoplatanus, Alnus glutinosa, Alnus incana, Fraxinus excelsior, Quercus cerris, Q. petraea, Q. robur, Q. delachampii, and Ulmus sp.
PMCID: PMC3755195  PMID: 23997626
Betula spp.; Birch; Chaga; Cinder conk; Inonotus obliquus; Sterile conks
18.  Seasonal dynamics of arboreal spider diversity in a temperate forest 
Ecology and Evolution  2012;2(4):768-777.
Measuring and estimating biodiversity patterns is a fundamental task of the scientist working to support conservation and inform management decisions. Most biodiversity studies in temperate regions were often carried out over a very short period of time (e.g., a single season) and it is often—at least tacitly—assumed that these short-term findings are representative of long-term general patterns. However, should the studied biodiversity pattern in fact contain significant temporal dynamics, perhaps leading to contradictory conclusions. Here, we studied the seasonal diversity dynamics of arboreal spider communities dwelling in 216 European beeches (Fagus sylvatica L.) to assess the spider community composition in the following seasons: two cold seasons (I: November 2005–January 2006; II: February–April) and two warm seasons (III: May–July; IV: August–October). We show that the usually measured diversity of the warm season community (IV: 58 estimated species) alone did not deliver a reliable image of the overall diversity present in these trees, and therefore, we recommend it should not be used for sampling protocols aimed at providing a full picture of a forest's biodiversity in the temperate zones. In particular, when the additional samplings of other seasons (I, II, III) were included, the estimated species richness nearly doubled (108). Community I possessed the lowest diversity and evenness due to the harsh winter conditions: this community was comprised of one dominant species together with several species low in abundance. Similarity was lowest (38.6%) between seasonal communities I and III, indicating a significant species turnover due to recolonization, so that community III had the highest diversity. Finally, using nonparametric estimators, we found that further sampling in late winter (February–April) is most needed to complete our inventory. Our study clearly demonstrates that seasonal dynamics of communities should be taken into account when studying biodiversity patterns of spiders, and probably forest arthropods in general.
PMCID: PMC3399199  PMID: 22837825
Araneae; canopy fogging; European beech; recolonization; species richness estimation; true diversity
19.  Differential Responses of Herbivores and Herbivory to Management in Temperate European Beech 
PLoS ONE  2014;9(8):e104876.
Forest management not only affects biodiversity but also might alter ecosystem processes mediated by the organisms, i.e. herbivory the removal of plant biomass by plant-eating insects and other arthropod groups. Aiming at revealing general relationships between forest management and herbivory we investigated aboveground arthropod herbivory in 105 plots dominated by European beech in three different regions in Germany in the sun-exposed canopy of mature beech trees and on beech saplings in the understorey. We separately assessed damage by different guilds of herbivores, i.e. chewing, sucking and scraping herbivores, gall-forming insects and mites, and leaf-mining insects. We asked whether herbivory differs among different forest management regimes (unmanaged, uneven-aged managed, even-aged managed) and among age-classes within even-aged forests. We further tested for consistency of relationships between regions, strata and herbivore guilds. On average, almost 80% of beech leaves showed herbivory damage, and about 6% of leaf area was consumed. Chewing damage was most common, whereas leaf sucking and scraping damage were very rare. Damage was generally greater in the canopy than in the understorey, in particular for chewing and scraping damage, and the occurrence of mines. There was little difference in herbivory among differently managed forests and the effects of management on damage differed among regions, strata and damage types. Covariates such as wood volume, tree density and plant diversity weakly influenced herbivory, and effects differed between herbivory types. We conclude that despite of the relatively low number of species attacking beech; arthropod herbivory on beech is generally high. We further conclude that responses of herbivory to forest management are multifaceted and environmental factors such as forest structure variables affecting in particular microclimatic conditions are more likely to explain the variability in herbivory among beech forest plots.
PMCID: PMC4132021  PMID: 25119984
20.  The Impact of Winter and Spring Temperatures on Temperate Tree Budburst Dates: Results from an Experimental Climate Manipulation 
PLoS ONE  2012;7(10):e47324.
Budburst phenology is a key driver of ecosystem structure and functioning, and it is sensitive to global change. Both cold winter temperatures (chilling) and spring warming (forcing) are important for budburst. Future climate warming is expected to have a contrasting effect on chilling and forcing, and subsequently to have a non-linear effect on budburst timing. To clarify the different effects of warming during chilling and forcing phases of budburst phenology in deciduous trees, (i) we conducted a temperature manipulation experiment, with separate winter and spring warming treatments on well irrigated and fertilized saplings of beech, birch and oak, and (ii) we analyzed the observations with five temperature-based budburst models (Thermal Time model, Parallel model, Sequential model, Alternating model, and Unified model). The results show that both winter warming and spring warming significantly advanced budburst date, with the combination of winter plus spring warming accelerating budburst most. As expected, all three species were more sensitive to spring warming than to winter warming. Although the different chilling requirement, the warming sensitivity was not significantly different among the studied species. Model evaluation showed that both one- and two- phase models (without and with chilling, respectively) are able to accurately predict budburst. For beech, the Sequential model reproduced budburst dates best. For oak and birch, both Sequential model and the Thermal Time model yielded good fit with the data but the latter was slightly better in case of high parameter uncertainty. However, for late-flushing species, the Sequential model is likely be the most appropriate to predict budburst data in a future warmer climate.
PMCID: PMC3468505  PMID: 23071786
21.  Phosphite Protects Fagus sylvatica Seedlings towards Phytophthora plurivora via Local Toxicity, Priming and Facilitation of Pathogen Recognition 
PLoS ONE  2014;9(1):e87860.
Phytophthora plurivora causes severe damage on Fagus sylvatica and is responsible for the extensive decline of European Beech throughout Europe. Unfortunately, no effective treatment against this disease is available. Phosphite (Phi) is known to protect plants against Phytophthora species; however, its mode of action towards P. plurivora is still unknown. To discover the effect of Phi on root infection, leaves were sprayed with Phi and roots were subsequently inoculated with P. plurivora zoospores. Seedling physiology, defense responses, colonization of root tissue by the pathogen and mortality were monitored. Additionally the Phi concentration in roots was quantified. Finally, the effect of Phi on mycelial growth and zoospore formation was recorded. Phi treatment was remarkably efficient in protecting beech against P. plurivora; all Phi treated plants survived infection. Phi treated and infected seedlings showed a strong up-regulation of several defense genes in jasmonate, salicylic acid and ethylene pathways. Moreover, all physiological parameters measured were comparable to control plants. The local Phi concentration detected in roots was high enough to inhibit pathogen growth. Phi treatment alone did not harm seedling physiology or induce defense responses. The up-regulation of defense genes could be explained either by priming or by facilitation of pathogen recognition of the host.
PMCID: PMC3905044  PMID: 24489973
22.  Fagus sylvatica trunk epicormics in relation to primary and secondary growth 
Annals of Botany  2012;110(5):995-1005.
Background and Aims
European beech epicormics have received far less attention than epicormics of other species, especially sessile oak. However, previous work on beech has demonstrated that there is a negative effect of radial growth on trunk sprouting, while more recent investigations on sessile oak proved a strong positive influence of the presence of epicormics. The aims of this study were, first, to make a general quantification of the epicormics present along beech stems and, secondly, to test the effects of both radial growth and epicormic frequency on sprouting.
In order to test the effect of radial growth, ten forked individuals were sampled, with a dominant and a dominated fork of almost equal length for every individual. To test the effects of primary growth and epicormic frequency, on the last 17 annual shoots of each fork arm, the number of axillary buds, shoot length, ring width profiles, epicormic shoots and other epicormics were carefully recorded.
Key Results
The distribution of annual shoot length, radial growth profiles and parallel frequencies of all epicormics are presented. The latter frequencies were parallel to the annual shoot lengths, nearly equivalent for both arms of each tree, and radial growth profiles included very narrow rings in the lowest annual shoots and even missing rings in the dominated arms alone. The location of the latent buds and the epicormics was mainly at branch base, while epicormic shoots, bud clusters and spheroblasts were present mainly in the lowest annual shoots investigated. Using a zero-inflated mixed model, sprouting was shown to depend positively on epicormic frequency and negatively on radial growth.
Support for a trade-off between cambial activity and sprouting is put forward. Sprouting mainly depends on the frequency of epicormics. Between- and within-tree variability of the epicormic composition in a given species may thus have fundamental and applied implications.
PMCID: PMC3448431  PMID: 22887022
European beech; Fagus sylvatica; epicormics; ontogeny; mixed ZIP models; sprouting; radial growth
23.  Simulating local adaptation to climate of forest trees with a Physio-Demo-Genetics model 
Evolutionary Applications  2014;7(4):453-467.
One challenge of evolutionary ecology is to predict the rate and mechanisms of population adaptation to environmental variations. The variations in most life history traits are shaped both by individual genotypic and by environmental variation. Forest trees exhibit high levels of genetic diversity, large population sizes, and gene flow, and they also show a high level of plasticity for life history traits. We developed a new Physio-Demo-Genetics model (denoted PDG) coupling (i) a physiological module simulating individual tree responses to the environment; (ii) a demographic module simulating tree survival, reproduction, and pollen and seed dispersal; and (iii) a quantitative genetics module controlling the heritability of key life history traits. We used this model to investigate the plastic and genetic components of the variations in the timing of budburst (TBB) along an elevational gradient of Fagus sylvatica (the European beech). We used a repeated 5 years climatic sequence to show that five generations of natural selection were sufficient to develop nonmonotonic genetic differentiation in the TBB along the local climatic gradient but also that plastic variation among different elevations and years was higher than genetic variation. PDG complements theoretical models and provides testable predictions to understand the adaptive potential of tree populations.
PMCID: PMC4001444  PMID: 24822080
budburst phenology; ecophysiology; European beech; Fagus sylvatica; individual-based model; quantitative genetics; quantitative trait loci
24.  Hitchhiking with forests: population genetics of the epiphytic lichen Lobaria pulmonaria in primeval and managed forests in southeastern Europe 
Ecology and Evolution  2012;2(9):2223-2240.
Availability of suitable trees is a primary determinant of range contractions and expansions of epiphytic species. However, switches between carrier tree species may blur co-phylogeographic patterns. We identified glacial refugia in southeastern Europe for the tree-colonizing lichen Lobaria pulmonaria, studied the importance of primeval forest reserves for the conservation of genetically diverse populations and analyzed differences in spatial genetic structure between primeval and managed forests with fungus-specific microsatellite markers. Populations belonged to either of two genepools or were admixed. Gene diversity was higher in primeval than in managed forests. At small distances up to 170 m, genotype diversity was lower in managed compared with primeval forests. We found significant associations between groups of tree species and two L. pulmonaria genepools, which may indicate “hitchhiking” of L. pulmonaria on forest communities during postglacial migration. Genepool B of L. pulmonaria was associated with European Beech (Fagus sylvatica) and we can hypothesize that genepool B survived the last glaciation associated within the refuge of European Beech on the Coastal and Central Dinarides. The allelic richness of genepool A was highest in the Alps, which is the evidence for a northern refuge of L. pulmonaria. Vicariant altitudinal distributions of the two genepools suggest intraspecific ecological differentiation.
PMCID: PMC3488673  PMID: 23139881
Conservation biology; lichen ecology; phylogeography; population genetics; primeval forests
25.  Comparison of pollen gene flow among four European beech (Fagus sylvatica L.) populations characterized by different management regimes 
Heredity  2011;108(3):322-331.
The study of the dispersal capability of a species can provide essential information for the management and conservation of its genetic variability. Comparison of gene flow rates among populations characterized by different management and evolutionary histories allows one to decipher the role of factors such as isolation and tree density on gene movements. We used two paternity analysis approaches and different strategies to handle the possible presence of genotyping errors to obtain robust estimates of pollen flow in four European beech (Fagus sylvatica L.) populations from Austria and France. In each country one of the two plots is located in an unmanaged forest; the other plots are managed with a shelterwood system and inside a colonization area (in Austria and France, respectively). The two paternity analysis approaches provided almost identical estimates of gene flow. In general, we found high pollen immigration (∼75% of pollen from outside), with the exception of the plot from a highly isolated forest remnant (∼50%). In the two unmanaged plots, the average within-population pollen dispersal distances (from 80 to 184 m) were higher than previously estimated for beech. From the comparison between the Austrian managed and unmanaged plots, that are only 500 m apart, we found no evidence that either gene flow or reproductive success distributions were significantly altered by forest management. The investigated phenotypic traits (crown area, height, diameter and flowering phenology) were not significantly related with male reproductive success. Shelterwood seems to have an effect on the distribution of within-population pollen dispersal distances. In the managed plot, pollen dispersal distances were shorter, possibly because adult tree density is three-fold (163 versus 57 trees per hectare) with respect to the unmanaged one.
PMCID: PMC3282401  PMID: 21897442
genetic diversity; paternity analysis; neighbourhood model; reproductive success; null alleles; nuclear microsatellites

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